taxonID	type	description	language	source
03BB8789FFB7FFAF2720D4F6FD65FF44.taxon	diagnosis	The three buthid genera Grosphus Simon, 1880, Neogrosphus Lourenço, 1995, and Teruelius gen. n. comprise a distinct assemblage of Madagascar buthids (= ‘ Grosphus ’ group) sharing the following set of characters: Carapace subrectangular, weakly trapezoidal or nearly parallel-sided, surface densely granular, carinae indistinct except for superciliary carinae; frontal region of carapace flat, not sloped towards anterior margin; median eyes large, median ocular tubercle prominent, located forward of the carapace centroid (Figs. 165 – 180); 5 pairs of lateral eyes (3 large, 2 small) (Figs. 227 – 230); chelicerae with typical buthid dentition on fixed and movable fingers (Vachon, 1963), two enlarged denticles on ventral surface of fixed finger (Figs. 231 – 238); sternum type 1, subtriangular; tergites granular, tergites I – VI with single, weak median carina, tergite VII with weak median carina and 2 pairs of strong lateral carinae; metasoma moderately elongate, segments I – III with 8 – 10 carinae, IV with 8 carinae, V with 3 – 5 carinae; telson vesicle bulbous, ovoid or elongate, with or without subaculear tubercle (Figs. 181 – 195); pectines with fulcra, 13 – 41 teeth, female with basal pectinal tooth dilated or elongated, lacking peg sensillae (Figs. 40 – 51, 196 – 210); hemispermatophore flagellum thicker at base, narrowed proximally, thickened distally (Figs. 52, 58, 60, 67, 71, 75, 78, 84); pedipalp chela elongate, smooth, carinae obsolete, surface typically with numerous short macrosetae (Figs. 21 – 24); finger dentition composed of 8 – 15 discrete linear rows of granules or denticles, each slightly oblique with proximal ends directed externally; rows either non-overlapping or slightly imbricated, proximal 3 granules in each row enlarged, 2 of these slightly displaced outwards as ‘ external accessory’ granules; series of large, dentate internal accessory granules present, offset from main rows; both chela fingers with enlarged apical teeth, 3 – 4 external subdistal granules; pedipalps sexually dimorphic, dentate margins of fingers weakly or strongly scalloped proximally in males, straight in females, manus of males broader than that of females; trichobothrial pattern orthobothriotaxic, type A (Vachon, 1974), with femur d 1 - d 3 - d 4 in α-configuration (Vachon, 1975), patella d 3 external to dorsomedian carina (Fet et al., 2005); patella em much closer to est and et, than to esb 1 and esb 2, with em - est - et usually forming a compact triad (Figs. 345, 481 a); chela manus with Eb 1 - Eb 2 angled distally, Eb 1 - Eb 2 - Eb 3 acute angle opening in proximal direction (γ-configuration) (Figs. 342, 478 a); chela with db in proximal half to middle of fixed finger; legs III – IV with tibial spurs (Figs. 211 – 226, 261 – 262, 318 – 319, 364 – 365, 414 – 417, 487 – 488, 514 – 515, 540 – 541, 578 – 579, 618 – 619), tarsi without bristle-combs.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFB7FFAF2720D4F6FD65FF44.taxon	discussion	REMARKS. In describing the first ‘ Grosphus ’ group species, Scorpio (Androctonus) madagascariensis, Gervais (1844: pl. XI, fig. 3) illustrated the carapace showing forward placement of the median eyes, and also accurately depicted five pairs of lateral eyes, now recognized to be the prevalent buthid configuration (Loria & Prendini, 2014; Yang et al., 2013). In spite of this, Fage (1929) incorrectly declared that Grosphus (sensu lato) only bore 3 pairs of lateral eyes, and Lourenço (1996 b) cited only 3 – 4 pairs. Moreover, only 3 pairs were described for: Grosphus ambre, G. darainensis, G. garciai, G. goudoti, G. halleuxi, G. hirtus, G. madagascariensis, G. makay, G. mandena, G. mayottensis, G. polskyi, G. rossii, G. simoni, G. rakotoariveloi, G. tavaratra, G. voahangyae, Teruelius ankarana, T. ankarafantsika, T. bemaraha, T. bicolor, T. bistriatus, T. eliseanneae, T. feti, T. ganzhorni, T. intertidalis, T. limbatus, T. magalieae, T. mahafaliensis, T. olgae, T. sabineae, T. waeberi (Lourenço, 1996 b, 1999, 2001 b, 2003 c, 2005, 2012 c, 2013 b, 2014; Lourenço & Goodman, 2006, 2009; Lourenço & Wilmé, 2015 a, 2015 b, 2016; Lourenço et al., 2004, 2007 a, 2009 b, 2016 c, 2017, 2018 b). We confirm here that 5 pairs are indeed present in all species that we have examined: G. garciai (= G. hirtus), G. goudoti, G. ‘ halleuxi ’, G. hirtus, G. sp. nr hirtus, G. madagascariensis, G. ‘ mandena ’, G. voahangyae, Neogrosphus griveaudi, Teruelius ankarafantsika, T. ankarana, T. annulatus, T. bistriatus, T. feti, T. flavopiceus, T. grandidieri, T. intertidalis, T. limbatus, T. mahafaliensis and T. olgae (e. g., Figs. 227 – 230). We found only a few individual deviations from the standard pattern, such as 2 large and 2 small ocelli, that we regarded as developmental anomalies. We predict that other ‘ Grosphus ’ group species will also comply with the 5 - eye pattern. Although undercounting of lateral eyes is perhaps attributable to overlooking of the smaller posterior and upper ocelli, 10 of the published 3 - eye counts post-date introduction of the 5 - eye model by Yang et al. (2013, coauthor Lourenço) and Loria & Prendini (2014). Paradoxically, Lourenço et al. (2007 a) claimed 3 lateral eyes in boilerplate descriptions of G. hirtus and G. polskyi, yet their figures clearly depict all 5 lateral eyes as being present in both species. Vachon (1969) correctly reported 5 “ nettement visibles ” lateral eyes, 3 large and 2 small, in both sexes of Neogrosphus griveaudi. Although 3 pairs were described for N. blanci and N. andrafiabe (Lourenço, 1996 b; Lourenço et al., 2015), we are skeptical that these counts are accurate.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFB9FFAA27A9D257FD18FF05.taxon	description	(Figs. 1 – 4, 9 – 12, 21 – 22, 25 – 43, 52 – 68, 86, 94 – 98, 106 – 125, 133 – 136, 145 – 149, 158 – 160, 165 – 169, 181 – 185, 196 – 200, 211 – 215, 227 – 228, 231 – 234, 239 – 386, 580 – 583, Tabs. 1 – 4)	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFB9FFAA27A9D257FD18FF05.taxon	type_taxon	TYPE SPECIES. Scorpio (Androctonus) madagascariensis Gervais, 1843.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFB9FFAA27A9D257FD18FF05.taxon	diagnosis	DIAGNOSIS. A member of the ‘ Grosphus ’ group differentiated as follows: medium-sized scorpions, adults ca. 25 – 75 mm in length; pedipalp finger granule rows 11 – 14 (Figs. 252, 286, 302, 330, 376), movable finger typically with 4 external subdistal granules; femur trichobothrium d 2 located on internal surface, or straddling dorsointernal carina (Figs. 9 – 12); chela manus with petite trichobothrium Eb 3 usually well separated from Eb 2, by more than half the distance between Eb 1 and Eb 2 (Figs. 21 – 22); manus trichobothrium V 2 roughly collinear with V 1 along chela axis or slightly displaced internally; lower pectinal tooth counts: ♂ 15 – 23, ♀ 12 – 19 (Figs. 28 – 31); basal pectinal tooth of females wide, oval to subrectangular, not distinctly longer than other teeth (Figs. 40 – 43, 196 – 200, 289); hemispermatophore capsule long or short, posterior lobe with long, lanceolate extension (Figs. 52 – 68); sternites with broad ovoid, elliptical or hemi-elliptical spiracles (Figs. 94 – 98); metasoma I with ventromedian carinae moderately to strongly crenulate or granulate (Figs. 122 – 125); telson with oval or bulbous vesicle, with or without subaculear tubercle in adults (Figs. 181 – 185); legs with ventral surface of telotarsus sparsely setose, with two rows of <20 short, setiform macrosetae (Figs. 133 – 137, 211 – 215, 259 – 262, 316 – 319, 362 – 365); telotarsus with dorsal terminal process of normal size; cuticle with weak UV fluorescence (Figs. 145 – 149).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFB9FFAA27A9D257FD18FF05.taxon	diagnosis	SUBORDINATE TAXA. Grosphus ambre Lourenço, Wilmé & Waeber, 2018 Grosphus darainensis Lourenço, Goodman & Ramilijaona, 2004 Grosphus goudoti Lourenço & Goodman, 2006 Grosphus hirtus Kraepelin, 1900 Grosphus madagascariensis (Gervais, 1843) Grosphus mayottensis Lourenço & Goodman, 2009 Grosphus polskyi Lourenço, Qi & Goodman, 2007 Grosphus rakotoariveloi Lourenço, Wilmé, Soarimalala & Waeber, 2017 Grosphus tavaratra Lourenço, Soarimalala & Goodman, 2009 Grosphus voahangyae Lourenço & Wilmé, 2015 See Tables 1 – 3 for diagnostic characters used to place the above taxa under Grosphus.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFB9FFAA27A9D257FD18FF05.taxon	discussion	REMARKS. We consider Grosphus paraphyletic and define two species groups distinguished by major differences in hemispermatophore capsule form: (i) ‘ madagascariensis ’ group: capsule elongate, monocarinate, with basal lobe located far proximal to base of flagellum (G. madagascariensis); (ii) ‘ hirtus ’ group: capsule short, carination variable, with basal lobe located distally near base of flagellum (G. goudoti, G. hirtus and G. voahangyae). Phylogenetic polarity of capsule form is unclear. Possible group affiliations of other species are suggested by some similarities in external characters, e. g.: (i) ‘ madagascariensis ’ group: elliptic spiracles, more elongate metasomal segments, maculation patterns weak or absent, subaculear tubercle small or absent; may include G. ambre, G. mayottensis and G. rakotoariveloi; (ii) ‘ hirtus ’ group: ovoid spiracles, more stout metasomal segments, stronger maculation patterns, subaculear tubercle more developed; may include G. polskyi, G. tavaratra. However, external characters can be misleading and definitive group assignment requires study of hemispermatophore capsules. For example, G. goudoti resembles species of the ‘ madagascariensis ’ group in external characters (metasoma slender, weak maculation, elliptic spiracles, lack of subaculear tubercle) but possesses a ‘ hirtus ’ group type of capsule. NEW SYNONYMIES. Grosphus halleuxi Lourenço, Wilmé, Soarimalala & Waeber, 2017 = Grosphus madagascariensis (Gervais, 1843), syn. n. Grosphus mandena Lourenço, 2005 = Grosphus madagascariensis (Gervais, 1843), syn. n. Grosphus simoni Lourenço, Goodman & Ramilijaona, 2004 = Grosphus madagascariensis (Gervais, 1843), syn. n. The single holotype male of G. madagascariensis used for description by Gervais (1843, 1844) is in poor condition after 176 years. It is disarticulated into several fragments: metasoma III – V + telson, metasoma I – II, hollowed carapace and tergites with most of coxosternal area and sternites III – VI missing, and 4 partial leg fragments (cf. https: // science. mnhn. fr / taxon / species / grosphus / madagascariensis). The type locality is given only as ‘ Madagascar’. Gervais (1844: pl. XI, figs. 1 – 3) published a color painting of the dorsal habitus, and drawings of two consecutive metasomal segments in lateral view (segments not specified, but possibly III – IV), showing enlarged spiniform granules on posterior dorsal carinae, and the carapace with median and lateral eyes. With only limited information available about the holotype, which has lost many body parts bearing key taxonomic characters, it is difficult to precisely pin down the identity of G. madagascariensis in relation to a group of several other currently-named similar taxa (i. e., G. darainensis, G. halleuxi, G. mandena, G. rakotoariveloi and G. simoni). Previous diagnoses of Kraepelin (1900), Fage (1929) and Lourenço (1996 b) listed some characters that differentiate G. madagascariensis from G. hirtus or Teruelius gen. n. Meristic characters were: PTC ♂ 18 – 20, ♀ 16 – 18, and pedipalp movable finger granule rows 12. Lourenço & Goodman (2006) suggested that Goudot, collector of the holotype, travelled in the north eastern region. They selected a male and female from Forêt de Plateau de Makira, in humid northeastern forest near Antongil Bay, as reference material for a redescription. The redescription is generic for the group, with few diagnostic characters: PTC ♂ 20, ♀ 15 – 16, pedipalp movable finger granule rows 13. Lateral eyes were incorrectly cited only as only 3 pairs, contradicting Gervais (1844). Grosphus simoni was described by Lourenço, et al. (2004) from two specimens: the holotype male from Forêt de Plateau de Makira, Forêt de Sahantaha, which is humid tropical forest in the northeast; and a paratype male from Station Forestière d’Ampijoroa, Ankarafantsika National Park, which is dry deciduous forest in the northwest. The differential diagnosis was brief: paler coloration, metasoma with strong granules and carinae, including several larger posterior spiniform granules on dorsal carinae on segments II – IV. Recently, Lourenço, et al. (2017) moved the paratype to a different species, G. rakotoariveloi, invalidating the original diagnosis of G. simoni based on both specimens. They revised the diagnosis of G. simoni as: moderately darker coloration, several larger posterior spiniform granules on dorsal carinae of metasoma II – IV, PTC ♂ 15 – 17, ♀ 14 – 15, pedipalp finger granule rows ♂ 11 – 12, ♀ 12 – 13, male chela with weak to moderate scalloping. Photographs were included for G. simoni specimens of both sexes from Forêt de Sahantaha (figs. 2 – 5), although the male was misidentified as a female, and the female misidentified as a male. The holotype male of G. simoni is well documented in high resolution images published on the FMNH website: https: // collections-zoology. fieldmuseum. org / catalogue / 963985. We studied FMNH materials (5 ♂, 1 ♀) from Andasibe determined as G. simoni. We further compared other materials, including a male determined by M. Vachon as G. madagascariensis (MHNG). We found no convincing diagnostic characters to support a distinction between G. simoni and G. madagascariensis. Diagnostic characters for G. simoni involve relatively minor differences in darker vs. lighter shades of color, differences in size of spiniform granules on dorsal metasomal segments, meristic differences of one or two pectine teeth with contiguous or overlapping ranges of PTC, and / or pedipalp finger granule row counts. These characters are subject to inter-population and geographic variation in many scorpion taxa. Allowing for typical genetic variation, the metasoma and telson of holotypes of G. simoni and G. madagascariensis do not differ significantly in carination, spination or morphometrics. In the absence of quantitative analysis showing discontinuous variation either in characters or morphometrics to support splitting into discrete species, we regard them as synonyms. Localities of G. simoni overlap or are sympatric with the distribution for G. madagascariensis in northeastern humid forests. The species G. rakotoariveloi has meristics (PTC ♂ 18 – 19, pedipalp granule rows ♂ 13 – 14) that also overlap or are contiguous with those of G. madagascariensis. However, the type (and only known) locality is in a different bioclimatic region with dry deciduous forest, disjunct from eastern humid forests. It has much lighter coloration and relatively wide pedipalp chelae. We provisionally list this species, until additional data are available. Grosphus halleuxi was described by Lourenço, et al. (2017) from a series of males from Torotorofotsy Forest, ca. 20 km NW of Moramanga, in a central humid forest area that is locally less humid than other eastern forests. The diagnostic characters for differentiating it from G. simoni were: darker coloration, smaller size of 55 mm, PTC ♂ 16 – 19, pedipalp granule rows ♂ 11 – 12, and weaker scalloping of pedipalp fingers. The meristic counts do not yield a differential diagnosis as they overlap those of G. simoni (= G. madagascariensis). We analyzed a series of near topotypic specimens (5 ♂, 6 ♀) from Moramanga and ca. 30 km E of Moramanga, whose males closely match photos of the G. halleuxi male holotype (Figs. 1 – 2, cf. Lourenço, et al., 2017: figs. 16 – 17). We obtained meristics: PTC ♂ 15 – 18, ♀ 13 – 15, pedipalp finger granule rows ♂ ♀ 12, which are not distinguishable from meristic ranges of G. madagascariensis. Other characters of darker or lighter shades, and degree of pedipalp finger scalloping can also be variable between populations. For example, in Figs. 1 & 3, a female from Moramanga area is darker than a male from the same area, showing that intensity of coloration varies within the same population, weakening this diagnostic character. G. halleuxi was diagnosed as “ much darker ”, but ‘ G. simoni ’ appears as dark, if not darker (Lourenço, et al., 2017: figs. 2 – 5 vs. 16 – 17). It was argued that G. halleuxi is a narrow-ranged species adapted to a less humid local microclimate. The existence of local microendemic taxa should be supported by strong diagnostic characters. Until such characters are defined, we regard this species as a local population of G. madagascariensis. Grosphus mandena was described by Lourenço (2005) from near Fort Dauphin, in the Mandena region of southeastern coastal rainforest. Differential diagnostic characters were: lighter coloration, weaker metasomal carination, one larger spiniform granule on metasoma II – IV, and a more granulated telson. Meristics were: PTC ♂ 19 – 20, ♀ 15 – 17, pedipalp granule rows ♂ 12 – 13. As discussed above, these characters fall within ranges of variation for G. madagascariensis, including local populations given other species names. We loaned and studied the male holotype and a female paratype from MHNG and found them to be indistinguishable from G. madagascariensis. We note that the female paratype is considerably darker than the male holotype. This shows that intensity of coloration varies even within the type population, and is not a reliable diagnostic character (Figs. 348 – 351, 580 – 583). Lourenço & Wilmé (2016: fig. 36) showed a nonoverlapping discontinuous transition, at latitude ca. 22 ° S between the northern range of G. madagascariensis and the southern range of G. mandena. The transition latitude does not correspond to any boundary between centers of endemism as defined by the watershed model of Wilmé et al. (2006). Lourenço et al., (2009 b) suggested that the disjunction is recent, due to extirpation of south littoral rainforest by humans. More robust diagnostic characters and analysis of clinal vs. discontinuous variation is necessary to delimit the southern populations as a distinctive species. Until such characters are defined, we regard this species as a southern population of G. madagascariensis. The synonymies of G. halleuxi and G. mandena with G. madagascariensis are further supported by their identical hemispermatophores, all of which have a unique, elongated capsule architecture with a proximal basal lobe (cf. Figs. 52 – 53, 56 – 57 vs. Figs. 54 – 55). Grosphus hirtus garciai Lourenço, 2001 = Grosphus hirtus Kraepelin, 1900, syn. n. Grosphus garciai was described by Lourenço (2001 b) from Station Forestière d’Ampijoroa, Ankarafantsika National Park, based on an adult male holotype and a juvenile, collected by García Herrero. In the diagnosis, it was differentiated from G. madagascariensis, a quite different species, by: smaller size, maculated light and dark pigmentation, pedipalp granule rows ♂ 13, weaker spiniform granules on pedipalp and metasoma, and weaker scalloping of pedipalp fingers. Curiously, it was not compared to G. hirtus, which bears a much greater similarity. Subsequently, Lourenço & Goodman (2006) redescribed G. hirtus based on material also from Station Forestière d’Ampijoroa, Ankarafantsika National Park, also collected by García Herrero. G. hirtus was separated from G. garciai by yellowish rather than reddish brown color, and larger size (40 – 50 mm vs. 32 mm). Subsequently, Lourenço & Wilmé, (2015 a) downgraded G. garciai to the status of subspecies, G. hirtus garciai, supposedly a microendemic taxon in a “ local isolated population ”. It is morphologically identical to, and differentiated from the nominotypical G. hirtus only by smaller size. The subspecies is known only from the type locality and the nominotypical G. hirtus also occurs in the same area (Lourenço & Goodman, 2006). The species G. hirtus is distributed more widely over northwest Madagascar (Lourenço & Wilmé, 2016: fig. 36). We loaned and studied the holotype of G. garciai, as well as male and female topotypes from FMNH. We confirmed that it is morphologically indistinguishable from G. hirtus (Figs. 263 – 305). We question whether an animal population of somewhat smaller average body size compared to closely neighboring conspecifics, but otherwise identical to them, merits subspecies status. Local size variations of species may be caused by varying environmental conditions that limit growth rates and development. The size differential is exaggerated by the reported body lengths, a measurement that can vary with mesosomal expansion: G. hirtus 40 – 50 mm (Lourenço & Goodman, 2006); G. h. garciai 28 – 32 mm (Lourenço & Wilmé, 2015 a). These numbers imply that G. hirtus is at least 25 % longer. A more reliable size comparison for morphometrically similar scorpions would use carapace length. For example, we measured carapaces of G. hirtus: ♂ 5.5 mm, ♀ 5.0 mm, vs. G. h. garciai: ♂ 5.2 mm, ♀ 4.4 mm, i. e., only 6 – 12 % longer. Lourenço et al. (2007 a) redescribed G. hirtus with body lengths of ♂ 34.3 mm, ♀ 31.8 mm, and carapace lengths ♂ 4.1 mm, ♀ 4.3 mm. These are small enough to overlap measurements for G. h. garciai. Considering this overlap, we regard G. h. garciai as a synonym of G. hirtus. Grosphus polskyi is another species that is quite similar to G. hirtus. It was diagnosed by having paler color, with weaker, more diffuse maculate patterns restricted to carapace and tergites, slightly more elongate metasoma segment I, weak spination on dorsal metasomal carinae, and a slightly larger subaculear tubercle. The only known record is the single male holotype from Mikea Forest near Toliara, on the southwestern coast. Although this is quite far south of the southern-most record of G. hirtus (Lourenço & Wilmé, 2016), records of the latter are sparse, so it is unclear if it represents a disjunction. We provisionally list this species, until it can be critically evaluated by more material and analysis of variation.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFAA27B2D214FC77FE41.taxon	description	(Figs. 25 – 39, 69 – 70, 87, 106 – 121, 606 – 619, Tabs. 1 – 4)	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFAA27B2D214FC77FE41.taxon	diagnosis	DIAGNOSIS. A member of the ‘ Grosphus ’ group differentiated as follows: small-sized scorpions, adults ca. 24 – 30 mm in length; pedipalp finger granule rows 8 – 9 (Fig. 612 – 613, 616), movable finger with no more than 3 external subdistal granules; femur trichobothrium d 2 located on dorsal, carinal or internal surface; chela manus with petite trichobothrium Eb 3 usually well separated from Eb 2, by more than half the distance between Eb 1 and Eb 2 (Figs. 25 – 27); manus trichobothrium V 2 strongly displaced internally relative to V 1; higher pectinal tooth counts: ♂ 27 – 31, ♀ 27 – 29 (Figs. 28 – 31); basal pectinal tooth of females wide, oval, only slightly longer than other teeth (Fig. 614); hemispermatophore capsule short, posterior lobe rounded, without lanceolate extension (Fig. 70); sternites with moderately narrow spiracles (Figs. 610, 614); metasoma I with ventromedian carinae moderate, finely granulate; telson with elongate vesicle, without subaculear tubercle; legs with ventral surface of telotarsus densely setose, with> 20 long, filiform setae (Figs. 618 – 619); telotarsus with dorsal terminal process very small; cuticle with strong UV fluorescence.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFAA27B2D214FC77FE41.taxon	discussion	SUBORDINATE TAXA. Neogrosphus andrafiabe Lourenço, Wilmé & Waeber, 2015 Neogrosphus blanci Lourenço, 1996 Neogrosphus griveaudi (Vachon, 1969)	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFAA27B2D214FC77FE41.taxon	discussion	REMARKS. Neogrosphus shares some primitive characters with Grosphus, and some derived characters with Teruelius gen. n. (summarized in Table 4). One interpretation of this is that Neogrosphus is descended from an intermediate stage in the evolution of Teruelius gen. n. from a Grosphus - like ancestor. Other characters, such as small size, internal displacement of V 2, reduced dorsal terminal process of telotarsus and elongated telson appear to be autapomorphies for the genus.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFB224B1D341FDFCF845.taxon	description	(Figs. 5 – 8, 13 – 20, 23 – 39, 44 – 51, 71 – 85, 90 – 93, 99 – 105, 106 – 121, 137 – 144, 150 – 160, 170 – 180, 186 – 195, 201 – 210, 216 – 226, 229 – 230, 235 – 238, 387 – 579, 584 – 605, Tabs. 1 – 4) http: // zoobank. org / urn: lsid: zoobank. org: act: 54 CB 8128 - BCFD- 4 B 1 F-A 947 - 153 C 7 CDD 5 B 83	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFB224B1D341FDFCF845.taxon	type_taxon	TYPE SPECIES. Buthus limbatus Pocock, 1889.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFB224B1D341FDFCF845.taxon	etymology	ETYMOLOGY. The generic epithet Teruelius (masculine) is a patronym honoring Rolando Teruel from Cuba in recognition of his many important contributions to the knowledge of scorpions.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFB224B1D341FDFCF845.taxon	diagnosis	DIAGNOSIS. A member of the ‘ Grosphus ’ group differentiated as follows: medium-sized to large-sized scorpions, adults ca. 35 – 120 mm in length; pedipalp finger granule rows 10 – 15 (Figs. 402, 431, 452, 485 – 486, 508, 521, 529, 560), movable finger typically with 4 external subdistal granules; femur trichobothrium d 2 straddling dorsointernal carina, or located on dorsal surface (Figs. 13 – 20); chela manus with petite trichobothrium Eb 3 near Eb 2, closer than half the distance between Eb 1 and Eb 2 (Figs. 23 – 24); manus trichobothrium V 2 roughly collinear with V 1 along chela axis or slightly displaced internally; higher pectinal tooth counts: ♂ 25 – 41, ♀ 24 – 35 (Figs. 28 – 31); basal pectinal tooth of females wide, with elongate, tapering distal extension, distinctly longer than other teeth (Figs. 44 – 51, 201 – 210, 411, 510, 526); hemispermatophore capsule short, carinate, posterior lobe rounded, without lanceolate extension (Figs. 71 – 85); sternites with narrow, slit-like spiracles (Figs. 99 – 105); metasoma I with ventromedian carinae moderately to weakly crenulate or smooth to obsolete (Figs. 126 – 132); telson with oval or bulbous vesicle, without subaculear tubercle in adults (Figs. 186 – 195); legs with ventral surface of telotarsus densely setose or scopulate, with broad, brush-like strips of> 20 long filiform macrosetae (Figs. 138 – 144, 216 – 226, 409 – 417, 487 – 490, 512 – 515, 538 – 541, 576 – 579); telotarsus with dorsal terminal process of normal size; cuticle with strong UV fluorescence (Figs. 150 – 157).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFB224B1D341FDFCF845.taxon	discussion	SUBORDINATE TAXA. Teruelius ankarafantsika (Lourenço, 2003) comb. n. Teruelius ankarana (Lourenço & Goodman, 2003) comb. n. Teruelius annulatus (Fage, 1929) comb. n. Teruelius bemaraha (Lourenço, Wilmé & Waeber, 2018) comb. n. Teruelius bicolor (Lourenço, 2012) comb. n. Teruelius bistriatus (Kraepelin, 1900) comb. n. Teruelius eliseanneae (Lourenço & Wilmé, 2016) comb. n. Teruelius feti (Lourenço, 1996) comb. n. Teruelius flavopiceus (Kraepelin, 1900) comb. n. Teruelius ganzhorni (Lourenço, Wilmé & Waeber, 2016) comb. n. Teruelius grandidieri (Kraepelin, 1900) comb. n. Teruelius intertidalis (Lourenço, 1999) comb. n. Teruelius limbatus (Pocock, 1889) comb. n. Teruelius magalieae (Lourenço, 2014) comb. n. (= T. mahafaliensis?) Teruelius mahafaliensis (Lourenço, Goodman & Ramilijaona, 2004) comb. n. Teruelius olgae (Lourenço, 2004) comb. n. Teruelius sabineae (Lourenço & Wilmé, 2016) comb. n. Teruelius waeberi (Lourenço & Wilmé, 2016) comb. n. See Tables 1 – 3 for diagnostic characters used to place the above species under Teruelius gen. n.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FFBCFFB224B1D341FDFCF845.taxon	discussion	REMARKS. Recognition of Teruelius gen. n. as a separate genus, distinct from Grosphus, necessitates revision of some previous concepts about taxonomy and biogeography of Grosphus. Lourenço et al. (2017) associated G. ‘ simoni ’ (= G. madagascariensis) with G. rakotoariveloi and G. ‘ halleuxi ’, and subsequently Lourenço et al. (2018 b) elaborated on a ‘ Grosphus simoni ’ group, treating it as a monophyletic unit of closely related “ sister ” species including the aforementioned three, plus G. ambre, G. bemaraha and G. mahafaliensis. A group diagnosis was not provided but, for G. bemaraha, mention was made of “ a number of features such as spiniform granules on the dorsal carinae of metasomal segments II-IV and on internal carinae of pedipalp femur and patella ”. We place G. bemaraha under Teruelius gen n., on the basis of narrow spiracles, dense tarsal setation and high pectinal tooth count (Table 3), and consider spiniform granules shared with G. simoni to be a homoplasy. G. bemaraha was claimed to be closer to G. rakotoariveloi, but these species belong to different genera. Lourenço et al. (2018 b: 74, fig. 1) included G. mahafaliensis in the ‘ simoni ’ group, perhaps due to similarities to G. bemaraha, noting in particular a high number of pectine teeth (but it lacks spiniform granules on metasomal carinae). In contrast, we find that T. mahafaliensis comb. n. is very far removed from G. ‘ simoni ’ (= G. madagascariensis), differing in all nine genus-level diagnostic characters, and in the fundamental architecture of the hemispermatophore capsule (cf. Fig. 84 vs. Figs. 52 – 57). Lourenço et al. (2018 b) discussed biogeographic hypotheses attempting to explain the distribution of the incongruous, polyphyletic ‘ Grosphus simoni ’ group. The very wide distribution of the group meant that it was “ adapted to humid, dry and subarid environments ”, and the two most disjunct species, G. bemaraha and G. mahafaliensis from western and southern localities, were speculated to “ belong to relict populations, which may have survived in humid refugia encountered in the sedimentary basins during the dry episodes of the paleoclimate oscillations ”. However, these two species are not closely related to the other four group members, but belong to Teruelius gen. n., whose ancestors may have already been adapted to dry environments. Although Pleistocene climatic fluctuations could be relevant for recent speciation events in Grosphus and Teruelius gen. n., the many correlated characters separating these two genera suggest a far earlier split, as in other taxa. For example, dated molecular phylogenies of Madagascar archaeid spiders (Wood et al., 2015), Brookesia and other chameleons (Tolley et al., 2013; Townsend et al., 2009) and Zonosaurus plated lizards (Blair et al., 2015) have revealed that the majority of divergences in these other endemic taxa are quite deep, occurring long before the advent of Pleistocene climate cycles. Species of Teruelius gen. n. can be loosely subdivided by size and coloration: large species, T. flavopiceus, T. ankarana, T. grandidieri and T. bicolor; species with patterns of dark stripes on tergites (‘ bistriatus ’ group of Lourenço & Wilmé, 2016): T. ankarafantsika, T. bistriatus, T. eliseanneae, T. feti, T. limbatus, T. sabineae and T. waeberi; species with almost uniform yellow, orange or brown tergites, and maybe darker metasoma IV or V: T. annulatus, T. bemaraha, T. ganzhorni, T. intertidalis, T. magalieae, T. mahafaliensis and T. olgae. These groupings have been used to construct species keys, in conjunction with some other characters including shapes of female basal pectine teeth (Fage, 1929; Lourenço, 2003 c, 2004 a, 2014; Lourenço et al., 2007 b; Vachon, 1969). Monophyly of these groupings remains to be tested. NEW SYNONYMIES. Grosphus makay Lourenço & Wilmé, 2015 = Teruelius feti (Lourenço, 1996) comb. n., syn. n. Grosphus feti was described by Lourenço (1996 b) from a juvenile male holotype, ostensibly collected from “ Prov. Tulear, Tanjon’ I Vohimena [= Cap Sainte Marie] Réserve spéciale, X. 1995 ” and deposited in FMNH. We loaned and studied the holotype and a second juvenile male labeled as “ ♂ paratype ” (which we concur is conspecific with the holotype). Until now, this species was only known from these two types. Associated with the type, we found locality labels (Fig. 459) that differ from the published type locality: “ MADAGASCAR: Province de Toliara, Fôret de Vohimena, 35 km SE Sakaraha, 17 - 24. i. 1996, MyrCE- 7541, 22 ° 41.0 ’ S 44 ° 49.8 ’ E, 780 m, S. M. Goodman 0000 011 031 FMNH- INS Grosphus feti Lourenço HOLOTYPE: det. 1996 ”. This locality is ca. 325 km roughly north of the published type locality at Cap Sainte Marie and the collection date of January 1996 is several months later. Either a labeling error occurred after description, or the published type locality is incorrect. Cap Sainte Marie is a biological study area and frequent source of scorpion materials (e. g., Lourenço & Wilmé, 2016) so data labels of specimens could have been confused. The FMNH label site is ca. 145 km south and slightly west of the type locality of Grosphus makay (Lourenço & Wilmé, 2015 b): “ Region Atsimo-Andrefana, ex Province of Toliara, Makay Mts., General Collection, dry-Forest on sandy soil, 12 / III / 2010 (B. L. Fisher et al.). BLF 25549. Female holotype (CAS) ”. These two localities have sandstone substrates, similar elevation and are located in the same general bioclimatic region. Comparison of the published habitus of the holotype adult female of T. makay and the holotype juvenile male of T. feti revealed very similar morphology and morphometrics even though sex and age differ. Most notably, their color patterns are identical in all details including: pattern of fuscosity on interocular triangle of carapace with pale cut-out behind lateral eyes; thin median line, precise fuscous banding patterns and transverse lateral striping on all tergites; darkly marked ventrolateral and ventromedian carinae on metasoma I – IV; fuscous patterns on metasoma V and telson; short fuscous strip on interno-proximal margin of pedipalp patella; leg femora with distal short, pale cut-outs on distal dark areas of prolateral surfaces and pale narrow lines on dorsal margins; and leg patellae with fuscous ventral margins on prolateral surfaces (compare Fig. 459 to fig. 13 of Lourenço & Wilmé, 2015 b). These very particular details of pigmentation pattern are not found in T. limbatus which was regarded as a closely related species (Figs. 5 – 8, 516 – 521). We also examined two adult females, near topotypes of G. makay from the Makay Mountains, that exhibited the same coloration patterns and morphometrics as T. feti (Fig. 459 a). We allowed for the fact that juveniles of pigmented scorpions usually display darker, more intense markings, and that in adults these color patterns are somewhat faded. Details of coloration pattern have been given high priority as characters for species-level taxonomy of Grosphus (Lourenço et al., 2009 b; Lourenço, 2014). We therefore consider Grosphus makay to be a junior synonym of T. feti, and the correct type locality of the latter to be that indicated on FMNH data labels. T. feti was never again collected from Cap Sainte Marie in over two decades of fieldwork since its description, although other scorpion species (e. g., T. sabineae) were discovered there. The adult male remains unknown. Our opinion could be verified by collection and analysis of topotypic adult and juvenile specimens from the FMNH locality. The description of T. feti (Lourenço, 1996 b: 14) noted the juvenile status of the holotype which is ca. 30 mm, but Lourenço (2014: 636) diagnosed the species as “ of small size with a total length of 30 to 40 mm ”. Small size cannot be a species diagnostic character if it is a property of the juvenile, and adult males are probably medium-sized, comparable to an adult female of G. makay, ca. 56 mm in body length. Our synonymy also implies that this is not a microendemic species of the upper Central Menabe, as suggested of Lourenço & Wilmé (2015 b). The T. feti emended type locality lies a significant distance south of the Makay mountains, in the Mangoky watershed (Wilmé et al., 2006). Grosphus rossii Lourenço, 2013 = Teruelius mahafaliensis (Lourenço, Goodman & Ramilijaona, 2004) comb. n., syn. n. Grosphus rossii was described by Lourenço (2013 b) from a single adult male holotype collected from “ Central region, NE Manandona, S of Antsirabe, in secondary growth forest, under log, 8 August 2004, W. R Lourenço ” and deposited in ZMUH. We loaned and studied the holotype and found that it was virtually identical in coloration, external morphological characters and morphometrics to Teruelius mahafaliensis (Lourenço, Goodman & Ramilijaona, 2004) comb. n. Our comparative materials of the latter included a male collected near the type locality of that species (Figs. 522 – 525), and determined material loaned from FMNH including two adult males. In his description, Lourenço (2013 b: 59) compared and contrasted G. rossii to T. limbatus, but not to T. mahafaliensis. A potential diagnostic difference is the pectinal tooth count (= 28) in G. rossii being lower than the range (35 – 40) reported for male T. mahafaliensis by Lourenço et al., (2007 b: 373, tab. III). However, we examined a male T. mahafaliensis collected near the species type locality with PTC of 29 – 33 (Figs. 524 – 525). The type locality of G. rossii on the central plateau at ca. 1400 m a. s. l., is in a cooler, more humid zone, quite far from the other records of T. mahafaliensis concentrated on the Mahafaly Plateau, a region of subarid thorn scrub along the southwest coast (ca. 120 m a. s. l.). It was suggested that G. rossii was evidence of microendemism. We take a more conservative position and interpret the very close morphologies of G. rossii and T. mahafaliensis as indicative of a eurytopic species with wider distribution. Broad elevation ranges of> 1400 m are known for some widely distributed scorpions that inhabit varied bioclimatic zones (e. g., Anuroctonus pococki Soleglad & Fet, 2004, 300 – 1850 m a. s. l., Soleglad & Fet, 2004; Bothriurus burmeisteri Kraepelin, 1894 and Brachistosternus weijenberghi (Thorell, 1876), 1000 – 3000 m a. s. l., Campón et al., 2014; Compsobuthus maindroni (Kraepelin, 1901), Hottentotta jayakari (Pocock, 1895), Nebo omanensis Francke, 1980 and Orthochirus glabrifrons (Kraepelin, 1903), 0 – 1850 m a. s. l., Lowe, 2010 c). The status of G. rossii should be reviewed when topotypic females are collected and their basal pectinal tooth compared to that of T. mahafaliensis, as this is a more reliable diagnostic character in Teruelius gen. n. Another potential synonym of T. mahafaliensis is T. magalieae (Lourenço, 2014). According to its description, the morphometrics, coloration, and meristics of T. magalieae are very close to those of T. mahafaliensis. The type locality of Cap Saint Marie (holotype male as only known specimen) lies on the southwestern coast in the same bioclimatic region as the latter species. Lourenço (2014: 633) did not compare T. magalieae to T. mahafaliensis, but claimed that the most closely related species was G. rossii, which we here synonymize under T. mahafaliensis. The diagnostic differences between T. magalieae and G. rossii are not compelling: (i) pectines with 36 vs. 28 teeth (a range of variation allowed here for T. mahafaliensis); (ii) pedipalp fingers with 12 – 13 vs. 12 – 12 granule rows (overlapping counts); and (iii) overall paler coloration (differences in color shade are not uncommon for different populations of a species inhabiting areas with different substrates). We provisionally list this species, until it can be critically evaluated by study of more material and analysis of variation. The female of T. magalieae is unknown, and the species might be better diagnosed if the female basal pectinal tooth were determined to be unique.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8DFF9B2461D66BFBE1FB8C.taxon	materials_examined	1 ♂ (holotype, Figs. 58 – 59, 88, 211, 239 – 262), Antsiranana Province, Forêt de Bobankota, Versant ouest, site No. 2, 11 km E of Daraina, 13 ° 13.414 ’ S 49 ° 45.586 ’ E, 350 – 550 m a. s. l., X. 2002 - III. 2003, leg. M. Raheriarisena & H. A. Rakotondravony (MHNG).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8CFF9A2722D193FDFCF9A4.taxon	materials_examined	1 ♂ (holotype of Grosphus garciai Lourenço, 2001, Figs. 66, 291 – 292, 294 – 305) 1 juv. (paratype, Figs. 213, 293), Majunga Province, Ankarafantsika Reserve, Ampijoroa, 16 ° 18 ’ 45.2 ” S 46 ° 48 ’ 54.2 ” E, 73 m a. s. l., VI. 2000, leg. García Herrero (MHNG); 1 ♀ (labeled Grosphus madagascariensis, Fig. 197), Majunga Province, Ankarafantsika Reserve, Forest Station Ampijoroa, Ampijoroa village, 16 ° 18 ’ 45.2 ” S 46 ° 48 ’ 54.2 ” E, 73 m a. s. l., VI. 2000, leg. García Herrero (MHNG); 1 ♂ (labeled Grosphus garciai), Majunga Province, Ankarafantsika Reserve, Forest Station Ampijoroa, 16 ° 18 ’ S 46 ° 48 ’ E, sand area of Paquypodium (= Pachypodium), 27. II. 2001 - 1. III. 2001, leg. García Herrero (MHNG); 4 ♂ 4 ♀ 1 juv. (Figs. 64, 196, 212, 263 – 290), southwestern region, inland zone between Ranohira and Llakaka, IX. 2004, leg. W. R. Lourenço (ZMUH); 1 ♂ (Figs. 11, 21, 96, 122, 134, 181), Antsiranana Province, Reserve Special d’Analamerana, Fôret d’Ankavanana, 15.8 km SE Anivorano- Nord, 12 ° 47.7 ’ S 49 ° 22.1 ’ E, 200 m a. s. l., 23. I. 2004, pitfall trap, in particularly disturbed mixed dry deciduous and humid forest, leg. S. M. Goodman, SMG # 14135 (FMNH 86976); 1 ♀ (Figs. 11, 42, 96, 146, 167, 182), Mahajanga Province, Forêt de Beanka, 18 ° 01 ’ 23 ” S 44 ° 30 ’ 08 ” E, 220 m a. s. l., slightly disturbed dry deciduous forest, leg. Z. H. Harimpitia, Z. H H- 032 (FMNH 3482761); 1 ♂ (Fig. 60, 62), N Antsiranana Province, Diego Suarez env., E of Ramena village, 12 ° 15 ’ 9.95 ” S 49 ° 21 ’ 31.05 ” E, ca. 50 m a. s. l., (FKCP, GLPC); 1 ♂ (Fig. 61), Mahajanga Province, Mahajamba riv., Ampatika env., 16 ° 08 ’ S 47 ° 15 ’ E, 2002 (FKCP, GLPC); 4 ♂ 3 ♀ 37 juvs. (Fig. 63), Majunga Province, Ankarafantsika Reserve, Forest Station Ampijoroa, “ Jardin Botanique A ”, 16 ° 18 ’ S 46 ° 48 ’ E, 24. - 24. II. 2001, leg. García Herrero (MHNG); 1 ♂ 1 ♀ (Figs. 65, 198), Toamasina Province, Forêt de Vohitaly, site F, 5 km SE village Anjiahely, 15 ° 26 ’ 58 ” S 49 ° 32 ’ 06 ” E, 540 – 680 m a. s. l., 28. XII. 2002, leg. V. Andrianjakarivelo (MHNG); 1 ♂ (Fig. 64), southwestern region, inland zone between Ranohira and Llakaka, IX. 2004, leg. W. R. Lourenço (ZMUH); 1 ♂ 1 ♀ (Figs. 21, 42, 97, 133, 149, 166), Mahajanga Province, SE d’Ampijoroa, 16 ° 19.4 ’ S 46 ° 48.4 ’ E, 160 m a. s. l., in dry deciduous forest on white sand, 20. IV. 2003, leg. S. M. Goodman, SMG # 13631, # 13632 pitfall 2 (FMNH ♂ 73434, ♀ 73436), det. as G. garciai.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8CFF9A2722D713FB4DFE65.taxon	materials_examined	3 ♂ (Fig. 56) 6 ♀ 2 juvs., ‘ Madagascar’, leg. Saussure, det. M. Vachon (MHNG); 11 ♂ 41 ♀ (Figs. 1 – 4, 9, 22, 41, 54, 94, 123, 135, 145, 159 – 160, 165, 183, 199, 214, 227, 231 – 234), Moramanga env., Toamasina, Anjiro, 18 ° 52 ’ S 47 ° 59 ’ E, 1995 (FKCP, GLPC); 1 ♂ (Figs. 52 – 53, 86), Andasibe, Marie Guest House, 18.94727 ° S 048.41782 ° E, No. 1197 (FKCP, GLPC); 1 ♂ (G. mandena holotype, Figs. 215, 348 – 349), 2 ♂ 1 ♀ 1 juv. (G. mandena paratypes, Figs. 55, 200, 350 – 351), Toliara Province, Mandena - Fort Dauphin, littoral forest 10 km north of Fort Dauphin, 6 - 12 / I / 1999, leg. J. - B. Ramanamanjato (MHNG); 1 ♂ 2 ♀ (Figs. 341 – 347), Moramanga env., 1997, (FKCP); 1 ♂ 1 ♀, Andasibe-Mantadia, Anamalazaotra forest (FKCP); 1 ♀, Fianarantsoa district, Ranomafana env., 21 ° 13 ’ S 47 ° 25 ’ E, 1995 (FKCP); 5 ♂ 1 ♀ (Figs. 10, 95, 124, 136, 148, 168, 184, 306 – 340), Toamasina Province, Ambalafary Forest, 14.5 km SW Andasibe, 19 ° 02 ’ 38 ” S 48 ° 20 ’ 55 ” E, 995 m a. s. l., 11. III. 2012, dense, humid lowland and montane forest, leg. V. Soarimalala VS- 2142 (FMNH 3482757), det. as G. simoni; 2 ♂ 3 ♀ 1 juv. (Fig. 57), Toamasina Province, Andasibe- Mantadia, near Andasibe, Camp Feon’ny Ala, 938 m a. s. l., 18 ° 56.836 ’ S 48 ° 25.063 ’ E, (FKCP, GLPC).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8CFF9A2463D333FB5BFF64.taxon	materials_examined	4 ♂ 1 ♀ (paratypes, Figs. 12, 43, 67 – 68, 89, 98, 125, 137, 147, 169, 185, 228, 352 – 386), Toamasina Province, region of Alaotra-Mangoro, Moramanga District, Analamy Forest, 10 km E Ambohimanarivo Village, 18 ° 48 ’ 20.8 ” S 48 ° 21 ’ 38.1 ” E, 1006 m a. s. l., 15 – 31. I. 2009, dense humid forest, leg. V. Soarimalala VS- 2142 (FMNH 2992958).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8CFF9A2463D233FA99FF85.taxon	materials_examined	1 ♂ 1 ♀ 2 juvs. (Figs. 69 – 70, 87, 606 – 619), Toliara Province, Tsimanampetsotsa National Park, Andranovao camp, 15 m a. s. l., 24 ° 01.505 ’ S 43 ° 44.306 ’ E (FKCP, GLPC).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8CFF992463D293FD7EFD07.taxon	materials_examined	1 ♀ (holotype, Figs. 219, 389 – 390, 394 – 403, 406 – 408, 410 – 415) 1 ♂ (paratype, Figs. 201, 220, 387 – 388, 391 – 393, 404 – 405, 409, 416 – 417), Majunga Province, Ankarafantsika Reserve, Forest Station Ampijoroa, 16 ° 18 ’ S 46 ° 48 ’ E, sand area of Paquypodium (= Pachypodium), 27. II. 2001 - 1. III. 2001, leg. García Herrero (MHNG); 1 ♀ 45 newborn (paratypes), Majunga Province, Ankarafantsika Reserve, Forest Station Ampijoroa, “ Jardin Botanique A ”, 16 ° 18 ’ S 46 ° 48 ’ E, 24. - 24. II. 2001, leg. García Herrero (MHNG); 1 ♂ (Figs. 13, 78 – 79, 93, 99, 138, 150, 170), Mahajanga Province, SE d’Ampijoroa, 16 ° 19.4 ’ S 46 ° 48.4 ’ E, 160 m a. s. l., in dry deciduous forest on white sand, 17. IV. 2003, leg. S. M. Goodman, SMG # 13610 pitfall (FMNH 73423); 1 ♀ (Figs. 13, 44, 78 – 79, 99, 127), Mahajanga Province, SE d’Ampijoroa, 16 ° 19.4 ’ S 46 ° 48.4 ’ E, 100 m a. s. l., in dry deciduous forest on white sand, 22. IV. 2003, leg. S. M. Goodman, SMG # 13639 pitfall 3 bucket 74 (FMNH 73425); 1 ♂ 1 ♀ (Figs. 15, 24, 45, 80, 99, 171, 186 – 187), Mahajanga Province, Réserve Forestière de l’Ankarafantsika, 5 km SSE Ampijoroa, 16 ° 20.3 ’ S 46 ° 47.6 ’ E, 160 m a. s. l., 4 – 7. II. 1997, slightly disturbed deciduous forest, pitfall, leg. S. M. Goodman (FMNH ♂ 73430, ♀ 73432), det. as G. bistriatus.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8CFF992463D293FD7EFD07.taxon	description	Teruelius ankarana (Lourenço & Goodman, 2003) comb. n.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8CFF992463D293FD7EFD07.taxon	materials_examined	1 ♂ 1 ♀ (Figs. 14, 46, 71 – 72, 90, 100, 128, 141, 154 – 155, 172, 188), Antsiranana Province, Reserve Special d’Analamerana, Forêt d’Ankavanana, 15.8 km SE Anivorano-Nord, 12 ° 47.7 ’ S 49 ° 22.1 E, 200 m a. s. l., 23. I. 2004, pitfall trap, in particularly disturbed mixed dry deciduous and humid forest, leg. S. M. Goodman, SMG # 14114 (FMNH 86978); 8 ♂ 10 ♀ 5 juvs. ♂ 3 juvs. ♀ (Figs. 100, 202, 216, 418 – 421), Antsiranana Province, Ankarana NP, Diego Suarez env., E of Ramena village, 12 ° 57 ’ 43.4 ” S 49 ° 07 ’ 13.48 ” E, 126 m a. s. l., (FKCP, GLPC); 2 ♂ (after 4 th ecdysis) 2 ♂ (after 5 th ecdysis) 2 ♀, 2011 (FKCP); 1 ♂, Mahajanga Province, Ankofia riv., Ambodimanga env. (Bora) (FKCP).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992463D751FAEEFAE7.taxon	materials_examined	1 ♀ (holotype, Figs. 622 – 623), Orange River Colony, Bethulie (BMNH No. 1905.3.30.45 - 54); 1 ♂ 1 ♀ (Figs. 106, 620 – 621, 624 – 628, 635 – 638, 641), no exact locality (CUPC).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992463D7B1FB59FB27.taxon	materials_examined	1 ♂ (Figs. 161 – 164), KwaZulu-Natal Ndumo – Shokwe, 30. III. 2017, 26.874930 ° S 32.210920 ° E, leg. P. Just, F. Šťáhlavský, V. Opatová, C. Haddad, R. Booysen, A. Gomez, J. Ruch & J. Schneider, JA 1418 (CUPC).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992463D6F1FC3DF447.taxon	materials_examined	1 juv. (Figs. 629 – 634, 639 – 640), Mpumalanga, God’s Window, 24.874719 ° S 30.890959 ° E, 6. IV. 2017, leg. F. Šťáhlavský (CUPC).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992722D011FD46FE47.taxon	materials_examined	4 ♂ 1 ♀ (Figs. 77, 203, 218, 422 – 432) 1 juv. (♂), Toliara Province, Tsimanampetsotsa National Park, Andranovao camp, 15 m a. s. l., 24 ° 01.505 ’ S 43 ° 44.306 ’ E, 2014 (FKCP, GLPC); 1 ♀, Isalo Mts, Ranohira near Tulear, 1998 (FKCP).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992722D351FEFEFEC6.taxon	materials_examined	1 ♂ 1 ♀ (syntypes, Figs. 204, 221, 433 – 458), Tullear, Makabo, 5. VII. 1900, ZMUH.	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992722D3D1FCB4FF27.taxon	materials_examined	2 juv ♂ (holotype, Figs. 459 – 472, paratype), Toliara Province, Fôret de Vohilema, 35 km SE Sakaraha, 22 ° 41.0 ’ S 44 ° 49.8 ’ E, 780 m a. s. l., 17 – 24. I. 1996, leg. S. M. Goodman (FMNH holotype 11031, paratype 11032); 2 ♀ (topotypes of G. makay, Figs. 210, 226, 459 a), Toliara Province, Makay Mts., (FKCP).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992722D2F1FF4AF987.taxon	materials_examined	1 ♂ (Figs. 16, 101, 173, 189, 230), Toliara Province, Parc National de Bemaraha, Ankidrodroa, 2.5 km NE Bekopaka, 19 ° 7.9 ’ S 44 ° 48.5 ’ E, 100 m a. s. l., 25. XI. 2001, secondary dry forest, leg. S. M. Goodman SMG 12489 (FMNH 73453); 1 ♀ (Figs. 16, 47, 101, 129, 143, 153, 174), Majunga, Melaky, Antsalova, Antsalova, Tsiandro, Bemaraha Plateau, Ambakoa forest, near Befanazava River, 18 ° 47.838 ’ S 44 ° 52.904 ’ E, 1400 ft a. s. l., 17. I. 2006, valley marsh, pitfall 3, bucket 7, leg. H. A. Rakotondravony, HER 02557 (FMNH 73428); 1 ♂ (Fig. 75 – 76, 92), No. 1196 (FKCP, GLPC); 1 ♂ 6 ♀ 2 ♂ juvs. 1 ♀ juv., Montagne d`Ambre 30 km south of Antseranana (FKCP); 1 ♂ 3 ♀, no exact locality data, 2011 (FKCP); 1 juv. (Figs. 597 – 599, with duplicated metasoma, dead during 2 nd ecdysis) (FKCP); 3 ♂ 6 ♀ 1 ♀ juv. (Figs. 205, 217, 473 – 490), N Antsiranana Province, Tamatave, Plateau von Antsirana, Diego Suarez env., E of Ramena village, 12 º 15 ’ 9.95 ” S 49 º 21 ’ 31.05 ” E, ca 50 m a. s. l. (FKCP).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992722D491FD4AFB27.taxon	materials_examined	1 ♂ (Figs. 17, 71, 73 – 74, 102, 130, 175, 190, 491 – 494), Tuléar, Atsimo-Andrefana, Morombe, Nosy Ambositra, Antevankira, Antevankira forest, near the Antsakabe River, 21 ° 56.753 ’ S 44 ° 02.781 ’ E, 130 – 160 ft, 3. II. 2007, riparian valley along the Antsakabe River, in a rotten tree trunk, leg. H. A. Rakotondravony HER 03685 (FMNH 73446); 1 ♀ (Figs. 17, 48, 102, 142, 151), Toliara Province, Maheleotse 124 c River Onilahy, 23 ° 31.600 ’ S 44 ° 05.366 ’ E, 68 m, leg. Achile Rasehmionna (FMNH); 3 ♂ 1 ♀ 3 juvs (Figs. 206, 223), Toliara Province, Ankotofotsy, No. 17, 2011 (FKCP); 1 ♂, Toliara Province, Tsimanampetsotsa, Mitoho camp, 10 m a. s. l., 24 ° 02.838 ’ S 43 ° 45.138 ’ E (FKCP).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992722D6F1FD8EF447.taxon	materials_examined	1 ♀ (holotype, Figs. 207, 222, 495 – 515), Toliara Province, 3.5 km north of Tulear, IV. 1998, leg. N. Lutzmann (ZMUH); 1 ♂ 1 ♀ (Fig. 85), No. 1485 (FKCP, GLPC).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992463D191FA76FDA7.taxon	materials_examined	8 ♂ 6 ♀ (Figs. 5 – 8, 18, 23, 49, 82, 83, 91, 103, 126, 139, 157, 159 – 160, 176, 192, 229, 235 – 238), Fianarantsoa Province, Forêt d’Ianasana, 7 km W Itremo, at source of Atsirakamhaity River, 20 ° 36.1 ’ S 46 ° 34.3 ’ E, 1630 m a. s. l., 6. II. 1999, under rocks in Tapia forest, leg. S. M. Goodman (FMNH 73449); 2 ♂ 7 ♀ (Fig. 208, 225, 516 – 521), Central region, south of Antsirabe, 2006 (FKCP); 1 ♂ 3 ♀, Central region, south ofAntsirabe, 2010 (FKCP).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992463D371FB2EFF87.taxon	materials_examined	1 ♂ (holotype of Grosphus rossii, Figs. 522 – 523, 536 – 541), Central Region, NE Manandona, S of Antsirabe, 8. IX. 2004, leg. W. R. Lourenço in secondary growth forest, under log (ZMUH); 2 ♂ 2 ♀ (Figs. 19, 50, 84, 104, 131, 140, 156, 177 – 178, 193), Toliara Province, 10.5 km SE Itampolo (village), 24 ° 44.2 ’ S 44 ° 01.39 ’ E, 120 m a. s. l., 20. II. 2005, pitfall trap in disturbed spiny bush on Mahafaly Plateau, leg. V. Soarimalala & S. M. Goodman (FMNH 73598); 2 ♂ 1 ♀ (Figs. 84, 209, 224, 524 – 535), Toliara Province, Zombitse-Vohibasia, Isoky forrest margin, 692 m a. s. l., 22 ° 41.012 ’ S 44 ° 51.835 ’ E, (FKCP, GLPC); 4 ♂ 1 ♀, Fianarantsoa Province, Isalo, Ananalava forest margin, Tanambao (Mandabe) vill. env., 724 m a. s. l., 22 ° 35.028 ’ S 45 ° 7.672 ’ E (FKCP).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
03BB8789FF8FFF992463D291FAC0F9A7.taxon	materials_examined	1 ♂ (paratype, Figs. 542 – 546, 552 – 563, 576 – 579), Toliara Province, Fôret des Mikeas, 9.5 km W Ankiloaka, 22 ° 46.7 ’ S 43 ° 31.4 ’ E, 16. III. 2003, leg. S. Goodman & V. Soarmalala (MHNG); 1 ♂ (Figs. 20, 81, 105, 179, 194), Toliara Province, 10.5 km SE Itampolo (village), 24 ° 44.2 ’ S 44 ° 01.39 ’ E, 120 m a. s. l., 19. II. 2005, pitfall trap in disturbed spiny bush on Mahafaly Plateau, leg. V. Soarimalala & S. M. Goodman SMG # 14539 (FMNH 86968); 1 ♀ (Figs. 20, 51, 105, 132, 144, 152, 180, 195, 547 – 551, 564 – 575), Toliara Province, Forêt des Milua, 19 km SW Tamotamo, 21 ° 52.0 ’ S 43 ° 39.6 ’ E, 70 m a. s. l., 23. III. 2003, found in pitfall trap 16, leg. V. Soarimalala # VS 376 (FMNH 73624); 1 ♂ (paratype), south region, Toliara Province, Fôret de Mikea, 7.5 km NE Tsifotsa, 22 ° 48.0 ’ S 43 ° 26. ’ E, 60 m a. s. l., 21 - 25. II. 2003, leg. S. M. Goodman, V. Soarimalala, hemispermatophore examined (MHNG).	en	Lowe, Graeme, Kovařík, František (2019): Review of Grosphus Simon 1880 with description of Teruelius gen n a new buthid genus from Madagascar (Scorpiones Buthidae). Euscorpius 281: 1-128, DOI: 10.18590/euscorpius.2019.vol2019.iss281.1, URL: https://doi.org/10.18590/euscorpius.2019.vol2019.iss281.1
